Lakes (II)

(I have had some computer issues over the last couple of weeks, which was the explanation for my brief blogging hiatus, but they should be resolved by now and as I’m already starting to fall quite a bit behind in terms of my intended coverage of the books I’ve read this year I hope to get rid of some of the backlog in the days to come.)

I have added some more observations from the second half of the book, as well as some related links, below.

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“[R]ecycling of old plant material is especially important in lakes, and one way to appreciate its significance is to measure the concentration of CO₂, an end product of decomposition, in the surface waters. This value is often above, sometimes well above, the value to be expected from equilibration of this gas with the overlying air, meaning that many lakes are net producers of CO₂ and that they emit this greenhouse gas to the atmosphere. How can that be? […] Lakes are not sealed microcosms that function as stand-alone entities; on the contrary, they are embedded in a landscape and are intimately coupled to their terrestrial surroundings. Organic materials are produced within the lake by the phytoplankton, photosynthetic cells that are suspended in the water and that fix CO₂, release oxygen (O₂), and produce biomass at the base of the aquatic food web. Photosynthesis also takes place by attached algae (the periphyton) and submerged water plants (aquatic macrophytes) that occur at the edge of the lake where enough sunlight reaches the bottom to allow their growth. But additionally, lakes are the downstream recipients of terrestrial runoff from their catchments […]. These continuous inputs include not only water, but also subsidies of plant and soil organic carbon that are washed into the lake via streams, rivers, groundwater, and overland flows. […] The organic carbon entering lakes from the catchment is referred to as ‘allochthonous’, meaning coming from the outside, and it tends to be relatively old […] In contrast, much younger organic carbon is available […] as a result of recent photosynthesis by the phytoplankton and littoral communities; this carbon is called ‘autochthonous’, meaning that it is produced within the lake.”

“It used to be thought that most of the dissolved organic matter (DOM) entering lakes, especially the coloured fraction, was unreactive and that it would transit the lake to ultimately leave unchanged at the outflow. However, many experiments and field observations have shown that this coloured material can be partially broken down by sunlight. These photochemical reactions result in the production of CO₂, and also the degradation of some of the organic polymers into smaller organic molecules; these in turn are used by bacteria and decomposed to CO₂. […] Most of the bacterial species in lakes are decomposers that convert organic matter into mineral end products […] This sunlight-driven chemistry begins in the rivers, and continues in the surface waters of the lake. Additional chemical and microbial reactions in the soil also break down organic materials and release CO₂ into the runoff and ground waters, further contributing to the high concentrations in lake water and its emission to the atmosphere. In algal-rich ‘eutrophic’ lakes there may be sufficient photosynthesis to cause the drawdown of CO₂ to concentrations below equilibrium with the air, resulting in the reverse flux of this gas, from the atmosphere into the surface waters.”

“There is a precarious balance in lakes between oxygen gains and losses, despite the seemingly limitless quantities in the overlying atmosphere. This balance can sometimes tip to deficits that send a lake into oxygen bankruptcy, with the O₂ mostly or even completely consumed. Waters that have O₂ concentrations below 2mg/L are referred to as ‘hypoxic’, and will be avoided by most fish species, while waters in which there is a complete absence of oxygen are called ‘anoxic’ and are mostly the domain for specialized, hardy microbes. […] In many temperate lakes, mixing in spring and again in autumn are the critical periods of re-oxygenation from the overlying atmosphere. In summer, however, the thermocline greatly slows down that oxygen transfer from air to deep water, and in cooler climates, winter ice-cover acts as another barrier to oxygenation. In both of these seasons, the oxygen absorbed into the water during earlier periods of mixing may be rapidly consumed, leading to anoxic conditions. Part of the reason that lakes are continuously on the brink of anoxia is that only limited quantities of oxygen can be stored in water because of its low solubility. The concentration of oxygen in the air is 209 millilitres per litre […], but cold water in equilibrium with the atmosphere contains only 9ml/L […]. This scarcity of oxygen worsens with increasing temperature (from 4°C to 30°C the solubility of oxygen falls by 43 per cent), and it is compounded by faster rates of bacterial decomposition in warmer waters and thus a higher respiratory demand for oxygen.”

“Lake microbiomes play multiple roles in food webs as producers, parasites, and consumers, and as steps into the animal food chain […]. These diverse communities of microbes additionally hold centre stage in the vital recycling of elements within the lake ecosystem […]. These biogeochemical processes are not simply of academic interest; they totally alter the nutritional value, mobility, and even toxicity of elements. For example, sulfate is the most oxidized and also most abundant form of sulfur in natural waters, and it is the ion taken up by phytoplankton and aquatic plants to meet their biochemical needs for this element. These photosynthetic organisms reduce the sulfate to organic sulfur compounds, and once they die and decompose, bacteria convert these compounds to the rotten-egg smelling gas, H₂S, which is toxic to most aquatic life. In anoxic waters and sediments, this effect is amplified by bacterial sulfate reducers that directly convert sulfate to H₂S. Fortunately another group of bacteria, sulfur oxidizers, can use H₂S as a chemical energy source, and in oxygenated waters they convert this reduced sulfur back to its benign, oxidized, sulfate form. […] [The] acid neutralizing capacity (or ‘alkalinity’) varies greatly among lakes. Many lakes in Europe, North America, and Asia have been dangerously shifted towards a low pH because they lacked sufficient carbonate to buffer the continuous input of acid rain that resulted from industrial pollution of the atmosphere. The acid conditions have negative effects on aquatic animals, including by causing a shift in aluminium to its more soluble and toxic form Al³⁺. Fortunately, these industrial emissions have been regulated and reduced in most of the developed world, although there are still legacy effects of acid rain that have resulted in a long-term depletion of carbonates and associated calcium in certain watersheds.”

“Rotifers, cladocerans, and copepods are all planktonic, that is their distribution is strongly affected by currents and mixing processes in the lake. However, they are also swimmers, and can regulate their depth in the water. For the smallest such as rotifers and copepods, this swimming ability is limited, but the larger zooplankton are able to swim over an impressive depth range during the twenty-four-hour ‘diel’ (i.e. light–dark) cycle. […] the cladocerans in Lake Geneva reside in the thermocline region and deep epilimnion during the day, and swim upwards by about 10m during the night, while cyclopoid copepods swim up by 60m, returning to the deep, dark, cold waters of the profundal zone during the day. Even greater distances up and down the water column are achieved by larger animals. The opossum shrimp, Mysis (up to 25mm in length) lives on the bottom of lakes during the day and in Lake Tahoe it swims hundreds of metres up into the surface waters, although not on moon-lit nights. In Lake Baikal, one of the main zooplankton species is the endemic amphipod, Macrohectopus branickii, which grows up to 38mm in size. It can form dense swarms at 100–200m depth during the day, but the populations then disperse and rise to the upper waters during the night. These nocturnal migrations connect the pelagic surface waters with the profundal zone in lake ecosystems, and are thought to be an adaptation towards avoiding visual predators, especially pelagic fish, during the day, while accessing food in the surface waters under the cover of nightfall. […] Although certain fish species remain within specific zones of the lake, there are others that swim among zones and access multiple habitats. […] This type of fish migration means that the different parts of the lake ecosystem are ecologically connected. For many fish species, moving between habitats extends all the way to the ocean. Anadromous fish migrate out of the lake and swim to the sea each year, and although this movement comes at considerable energetic cost, it has the advantage of access to rich marine food sources, while allowing the young to be raised in the freshwater environment with less exposure to predators. […] With the converse migration pattern, catadromous fish live in freshwater and spawn in the sea.”

“Invasive species that are the most successful and do the most damage once they enter a lake have a number of features in common: fast growth rates, broad tolerances, the capacity to thrive under high population densities, and an ability to disperse and colonize that is enhanced by human activities. Zebra mussels (Dreissena polymorpha) get top marks in each of these categories, and they have proven to be a troublesome invader in many parts of the world. […] A single Zebra mussel can produce up to one million eggs over the course of a spawning season, and these hatch into readily dispersed larvae (‘veligers’), that are free-swimming for up to a month. The adults can achieve densities up to hundreds of thousands per square metre, and their prolific growth within water pipes has been a serious problem for the cooling systems of nuclear and thermal power stations, and for the intake pipes of drinking water plants. A single Zebra mussel can filter a litre a day, and they have the capacity to completely strip the water of bacteria and protists. In Lake Erie, the water clarity doubled and diatoms declined by 80–90 per cent soon after the invasion of Zebra mussels, with a concomitant decline in zooplankton, and potential impacts on planktivorous fish. The invasion of this species can shift a lake from dominance of the pelagic to the benthic food web, but at the expense of native unionid clams on the bottom that can become smothered in Zebra mussels. Their efficient filtering capacity may also cause a regime shift in primary producers, from turbid waters with high concentrations of phytoplankton to a clearer lake ecosystem state in which benthic water plants dominate.”

“One of the many distinguishing features of H₂O is its unusually high dielectric constant, meaning that it is a strongly polar solvent with positive and negative charges that can stabilize ions brought into solution. This dielectric property results from the asymmetrical electron cloud over the molecule […] and it gives liquid water the ability to leach minerals from rocks and soils as it passes through the ground, and to maintain these salts in solution, even at high concentrations. Collectively, these dissolved minerals produce the salinity of the water […] Sea water is around 35ppt, and its salinity is mainly due to the positively charged ions sodium (Na⁺), potassium (K⁺), magnesium (Mg²⁺), and calcium (Ca²⁺), and the negatively charged ions chloride (Cl^–), sulfate (SO₄^2-), and carbonate CO₃^2-). These solutes, collectively called the ‘major ions’, conduct electrons, and therefore a simple way to track salinity is to measure the electrical conductance of the water between two electrodes set a known distance apart. Lake and ocean scientists now routinely take profiles of salinity and temperature with a CTD: a submersible instrument that records conductance, temperature, and depth many times per second as it is lowered on a rope or wire down the water column. Conductance is measured in Siemens (or microSiemens (µS), given the low salt concentrations in freshwater lakes), and adjusted to a standard temperature of 25°C to give specific conductivity in µS/cm. All freshwater lakes contain dissolved minerals, with specific conductivities in the range 50–500µS/cm, while salt water lakes have values that can exceed sea water (about 50,000µS/cm), and are the habitats for extreme microbes”.

“The World Register of Dams currently lists 58,519 ‘large dams’, defined as those with a dam wall of 15m or higher; these collectively store 16,120km³ of water, equivalent to 213 years of flow of Niagara Falls on the USA–Canada border. […] Around a hundred large dam projects are in advanced planning or construction in Africa […]. More than 300 dams are planned or under construction in the Amazon Basin of South America […]. Reservoirs have a number of distinguishing features relative to natural lakes. First, the shape (‘morphometry’) of their basins is rarely circular or oval, but instead is often dendritic, with a tree-like main stem and branches ramifying out into the submerged river valleys. Second, reservoirs typically have a high catchment area to lake area ratio, again reflecting their riverine origins. For natural lakes, this ratio is relatively low […] These proportionately large catchments mean that reservoirs have short water residence times, and water quality is much better than might be the case in the absence of this rapid flushing. Nonetheless, noxious algal blooms can develop and accumulate in isolated bays and side-arms, and downstream next to the dam itself. Reservoirs typically experience water level fluctuations that are much larger and more rapid than in natural lakes, and this limits the development of littoral plants and animals. Another distinguishing feature of reservoirs is that they often show a longitudinal gradient of conditions. Upstream, the river section contains water that is flowing, turbulent, and well mixed; this then passes through a transition zone into the lake section up to the dam, which is often the deepest part of the lake and may be stratified and clearer due to decantation of land-derived particles. In some reservoirs, the water outflow is situated near the base of the dam within the hypolimnion, and this reduces the extent of oxygen depletion and nutrient build-up, while also providing cool water for fish and other animal communities below the dam. There is increasing attention being given to careful regulation of the timing and magnitude of dam outflows to maintain these downstream ecosystems. […] The downstream effects of dams continue out into the sea, with the retention of sediments and nutrients in the reservoir leaving less available for export to marine food webs. This reduction can also lead to changes in shorelines, with a retreat of the coastal delta and intrusion of seawater because natural erosion processes can no longer be offset by resupply of sediments from upstream.”

“One of the most serious threats facing lakes throughout the world is the proliferation of algae and water plants caused by eutrophication, the overfertilization of waters with nutrients from human activities. […] Nutrient enrichment occurs both from ‘point sources’ of effluent discharged via pipes into the receiving waters, and ‘nonpoint sources’ such the runoff from roads and parking areas, agricultural lands, septic tank drainage fields, and terrain cleared of its nutrient- and water-absorbing vegetation. By the 1970s, even many of the world’s larger lakes had begun to show worrying signs of deterioration from these sources of increasing enrichment. […] A sharp drop in water clarity is often among the first signs of eutrophication, although in forested areas this effect may be masked for many years by the greater absorption of light by the coloured organic materials that are dissolved within the lake water. A drop in oxygen levels in the bottom waters during stratification is another telltale indicator of eutrophication, with the eventual fall to oxygen-free (anoxic) conditions in these lower strata of the lake. However, the most striking impact with greatest effect on ecosystem services is the production of harmful algal blooms (HABs), specifically by cyanobacteria. In eutrophic, temperate latitude waters, four genera of bloom-forming cyanobacteria are the usual offenders […]. These may occur alone or in combination, and although each has its own idiosyncratic size, shape, and lifestyle, they have a number of impressive biological features in common. First and foremost, their cells are typically full of hydrophobic protein cases that exclude water and trap gases. These honeycombs of gas-filled chambers, called ‘gas vesicles’, reduce the density of the cells, allowing them to float up to the surface where there is light available for growth. Put a drop of water from an algal bloom under a microscope and it will be immediately apparent that the individual cells are extremely small, and that the bloom itself is composed of billions of cells per litre of lake water.”

“During the day, the [algal] cells capture sunlight and produce sugars by photosynthesis; this increases their density, eventually to the point where they are heavier than the surrounding water and sink to more nutrient-rich conditions at depth in the water column or at the sediment surface. These sugars are depleted by cellular respiration, and this loss of ballast eventually results in cells becoming less dense than water and floating again towards the surface. This alternation of sinking and floating can result in large fluctuations in surface blooms over the twenty-four-hour cycle. The accumulation of bloom-forming cyanobacteria at the surface gives rise to surface scums that then can be blown into bays and washed up onto beaches. These dense populations of colonies in the water column, and especially at the surface, can shade out bottom-dwelling water plants, as well as greatly reduce the amount of light for other phytoplankton species. The resultant ‘cyanobacterial dominance’ and loss of algal species diversity has negative implications for the aquatic food web […] This negative impact on the food web may be compounded by the final collapse of the bloom and its decomposition, resulting in a major drawdown of oxygen. […] Bloom-forming cyanobacteria are especially troublesome for the management of drinking water supplies. First, there is the overproduction of biomass, which results in a massive load of algal particles that can exceed the filtration capacity of a water treatment plant […]. Second, there is an impact on the taste of the water. […] The third and most serious impact of cyanobacteria is that some of their secondary compounds are highly toxic. […] phosphorus is the key nutrient limiting bloom development, and efforts to preserve and rehabilitate freshwaters should pay specific attention to controlling the input of phosphorus via point and nonpoint discharges to lakes.”

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Ultramicrobacteria.
The viral shunt in marine foodwebs.
Proteobacteria. Alphaproteobacteria. Betaproteobacteria. Gammaproteobacteria.
Mixotroph.
Carbon cycle. Nitrogen cycle. Ammonification. Anammox. Comammox.
Methanotroph.
Phosphorus cycle.
Littoral zone. Limnetic zone. Profundal zone. Benthic zone. Benthos.
Phytoplankton. Diatom. Picoeukaryote. Flagellates. Cyanobacteria.
Trophic state (-index).
Amphipoda. Rotifer. Cladocera. Copepod. Daphnia.
Redfield ratio.
δ15N.
Thermistor.
Extremophile. Halophile. Psychrophile. Acidophile.
Caspian Sea. Endorheic basin. Mono Lake.
Alpine lake.
Meromictic lake.
Subglacial lake. Lake Vostock.
Thermus aquaticus. Taq polymerase.
Lake Monoun.
Microcystin. Anatoxin-a.